Bioelectromagnetism Exercise #2 – Answers TAMPERE UNIVERSITY OF TECHNOLOGY Ragnar Granit...

Bioelectromagnetism Exercise #2 – Answers

TAMPERE UNIVERSITY OF TECHNOLOGYRagnar Granit Institute

Bioelectromagnetism Exercise 2


Q1: Characteristic Length and Time Constant

The intracellular resistance of a nerve cell is 8.2*106 Ω/cm (ri). Resistance of the cell membrane is 1.5*104 Ωcm and capacitance 12 nF/cm (cm). Calculate the characteristic length and time constant of the axon. (start from the general cable equation to see how the time constant is derived)



Review of terminology

Intracellular resistance & resistivity– Let R be the total resistance in axial direction [Ω]– > resistance per length

ri = R / l [Ω/m]

– > resistivity R=ρl/A

-> ρ = RA / l = ri * A [Ω m]

Extracellular resistance & resistivity

ro = R / l [Ω/m]

ρ = RA / l = ro * A [Ω m]

R

Q1: Characteristic Length and Time Constant - terminology

l

r

ri




Cell membrane resistance & resistivity– Let R be the total radial resistance [Ω]– > resistance in axial direction (as a function of the length of the membrane)

rm = R * l [Ω m]

– resistance is inversely proportional to the length of the membrane

– > resistivity ρm = RAm / lm = R (2 r l) / d [Ω m]

d = thickness of the membranel = length of the cell

– resistance per area Rm = R * A = R (2 r l) = rm * 2 r [Ω m2]

l

r

rm/l

r

d




Membrane capacitance– C total capacitance [F] (radial)

– > Capacitance per length cm = C / l [F/cm]

– > Capacitance per area Cm = C / A = C / ( 2 r * l) = cm / (2 r) [F/cm2]

l

r dR C




General cable equation describes passive function of a cell (subtreshold im) – 1-D propagation (along x-axis)

– V’ = Vm – Vr - deviation from RMP

– equivalent circuit2

2

' ( ) ' (3.41...3.45)i oi o m

m

r rV r r i Vrx

l

r

x




…

General solution of this equation is

boundary conditions: V’(x=0)=V’, V’(x=)=0

λ = characteristic length/length constant– describes spreading along the cell axis

– think: rm up -> λ up

2

2

' ' (3.41...3.45)i o

m

r rV Vrx

'x x

V Ae Be

0

m

i

r

r r




since ri >> ro =>

Time constant = rm*cm

– measure to reach steady-state

( 3.48)m eq

i

r

r

4

6

1.5*100.04277 428

8.2*10 /

cmcm m

cm

4 91.5*10 *12*10 / 180cm F cm s

0

m

i

r

r r



Q2: Strength-Duration Curve

The rheobasic current of the nerve cell in the previous exercise is 2 mA.a) What is the strength-duration equation of the cell. How long will it take to reach the stimulus threshold with a 2.5 mA stimulus current. What is the chronaxy of the cell?b) Determine the propagation speed of an action pulse if the cell diameter is 100 µm and coefficient K = 10.47 1/ms in propagation equation

ρ is the intracellular resistivity.

Definitions– Rheobase: smallest current, that generates an action

impulse

– Chronaxy: time, that is needed to generate action impulse with 2*Irh

K

2m

r

C




Definitions– impulse response of the membrane (radial direction only)

Vm

Is

Time

(3.56)/' (1 )s mtV I R e




RheobaseVth = membrane potential, that can generate action impulse

when t=:

== Rheobase

->

ChronaxyIs = 2 * Irh => t = * ln2 = 125 µs

/

'

(1 )s t

m

VIR e

/(1 )th th

s rh t

mm

V VI I

RR e

/

1/1 *ln(1 )

1

rh rhs t

rhs

s

I ItI e t

IIe

I

1180μs*ln 2902

(1 )2.5

s

0




Propagation speed

whereK = 10.47 1/msd = 100*10-6 mρ = intracellular resistivity

ρ = RA/l =Ri*ARi = R/l=8.2*106 Ω/cmρ = Ri * r2 = 8.2*106 Ω/cm * (5000*10-6cm)2 = 644 ΩcmCm = cm/(2r) = 12nF/cm / (2*5000*10-6cm) = 0.382 µF/cm2

=> 327 cm/s

empirical (eq. 4.33):

K

2m

r

C

r d



Q3: Sodium Conductance

Derive the equation of sodium conductance in voltage clamp measurements (with chemical clamping) using the Hodgkin-Huxley model.

Hodgkin-Huxley modelTransmembrane current equation

( ) ( ) ( )mm m m Na Na m K K m L L

dVI C V V G V V G V V G

dt

This is eq. 4.10 in the Bioelectromagnetism book




Hodgkin-Huxley model equations…




Transmembrane current

Voltage Clamp– no IC

Chemical Clamp– no INa

=>

Sodium current:

( ) ( ) ( )mm m m Na Na m K K m L L

dVI C V V G V V G V V G

dt

' ( ) ( )m m K K m L LI V V G V V G

' ' ( )Na m m m Na NaI I I V V G

'm mNa

m Na

I IGV V



Q4:Value of GNa

Cell membrane was studied with the voltage clamp measurement with a 56 mV positive voltage step. 2.5 ms after the step the membrane current is 0.6 mA/cm2. When the sodium current was blocked with pharmaceutical the current was 1 mA/cm2 (again, t = 2.5 ms after the step). Also, it was observed that the flow of sodium ions could be stopped with 117 mV increase in resting membrane potential. What is the sodium ion conductance GNa (stimulation 56 mV, 2.5 ms)?



Q4:Value of GNa

Voltage Clamp = no IC Two cases (56 mV voltage step)

1. no chemical clamping (t=2.5 ms): Im = 0.6 mA/cm2

2. chemical clamping (t=2.5 ms): I’m = 1.0 mA/cm2

I’m = IK (+ICl)

Im = IK + INa (+ICl)

=> INa = Im – I’m

INa can be blocked with 117 mV voltage step

VNa = Vr + 117 mV

Vm = Vr + 56mV



Q4:Value of GNa

GNa (56 mV, 2.5 ms) =

=> 65.6 S/m2

2

'

( 56 mV) - ( 117 mV)

0.6-1.0mA/cm(56 - 117) mV

m m

r r

i i

V V

'm mNa

m Na

I IGV V



Q5: BSM

The body surface ECG is measured using 26 to 256 electrodes. Figure 1 represents voltages of a normal body surface ECG measured at the end of a QRS complex. What can you say about the nature of the source according to this map?

Figure 1. Anterior body surface map (BSM).



Q5: BSM

Zero potential + -

dipolar field eq. dipole I source not normal BSM?

- inverted?

+

-

Bioelectromagnetism Exercise #2 – Answers TAMPERE UNIVERSITY OF TECHNOLOGY Ragnar Granit...

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